Inferring the biogeographic origins of inter‐continental disjunct endemics using a Bayes‐DIVA approach

The arcto‐Tertiary relictual flora is comprised of many genera that occur non‐contiguously in the temperate zones of eastern Asia, Europe, eastern North America, and western North America. Within each distributional area, species are typically endemic and may thus be widely separated from closely related species within the other areas. It is widely accepted that this common pattern of distribution resulted from of the fragmentation of a once more‐continuous arcto‐Tertiary forest. The historical biogeographic events leading to the present‐day disjunction have often been investigated using a phylogenetic approach. Limitations to these previous studies have included phylogenetic uncertainty and uncertainty in ancestral range reconstructions. However, the recently described Bayes‐DIVA method handles both types of uncertainty. Thus, we used Bayes‐DIVA analysis to reconstruct the stem lineage distributions for 185 endemic lineages from 23 disjunct genera representing 17 vascular plant families. In particular, we asked whether endemic lineages within each of the four distributional areas more often evolved from (1) widespread ancestors, (2) ancestors dispersed from other areas, or (3) endemic ancestors. We also considered which of these three biogeographic mechanisms may best explain the origins of arcto‐Tertiary disjunct endemics in the neotropics. Our results show that eastern Asian endemics more often evolved from endemic ancestors compared to endemics in Europe and eastern and western North America. Present‐day endemic lineages in the latter areas more often arose from widespread ancestors. Our results also provide anecdotal evidence for the importance of dispersal in the biogeographic origins of arcto‐Tertiary species endemic in the neotropics.     

Intercontinental disjunctions between eastern Asia and western North America in vascular plants highlight the biogeographic importance of the Bering land bridge from late Cretaceous to Neogene

This review shows a close biogeographic connection between eastern Asia and western North America from the late Cretaceous to the late Neogene in major lineages of vascular plants (flowering plants, gymnosperms, ferns and lycophytes). Of the eastern Asian–North American disjuncts, conifers exhibit a high proportion of disjuncts between eastern Asia and western North America. Several lineages of ferns also show a recent disjunct pattern in the two areas. In flowering plants, the pattern is commonly shown in temperate elements between northeastern Asia and northwestern North America, as well as elements of the relict boreotropical and Neogene mesophytic and coniferous floras. The many cases of intercontinental biogeographic disjunctions between eastern Asia and western North America in plants supported by recent phylogenetic analyses highlight the importance of the Bering land bridge and/or the plant migrations across the Beringian region from the late Cretaceous to the late Neogene, especially during the Miocene. The Beringian region has permitted the filtering and migration of certain plant taxa since the Pliocene after the opening of the Bering Strait, as many conspecific taxa or closely related species occur on both sides of Beringia.     

Harlequin duck Histrionicus histrionicus population structure in eastern Nearctic

During May 1996 and April 1997, eight harlequin duck males were captured and fitted with satellite transmitters while migrating along the shores of Forillon National Park, Québec, Canada. Another 17 males were equipped with satellite transmitters in river systems of eastern Hudson Bay, Ungava Bay and northern Labrador in June 1997 and 1998. Our objectives were to determine relationships between breeding, moulting and wintering areas, and to determine whether distinct population segments existed among harlequin ducks in eastern North America. All birds tracked from Forillon migrated to Labrador. Moulting areas were identified for six birds. Forillon males were followed to the eastern North American major wintering site in Maine. Males captured in northern Québec and Labrador migrated to moult and winter in south-western Greenland. Our data suggest the presence of two demographically distinct population segments in eastern North America, perhaps originating from the Pleistocene glacial refuge in western Greenland and south of the Laurentide ice sheet in eastern Canada or United States.     

Lack of genetic differentiation between monarch butterflies with divergent migration destinations

Monarch butterflies are best known for their spectacular annual migration from eastern North America to Mexico. Monarchs also occur in the North American states west of the Rocky Mountains, from where they fly shorter distances to the California Coast. Whether eastern and western North American monarchs form one genetic population or are genetically differentiated remains hotly debated, and resolution of this debate is essential to understand monarch migration patterns and to protect this iconic insect species. We studied the genetic structure of North American migratory monarch populations, as well as nonmigratory populations in Hawaii and New Zealand. Our results show that eastern and western migratory monarchs form one admixed population and that monarchs from Hawaii and New Zealand have genetically diverged from North American butterflies. These findings suggest that eastern and western monarch butterflies maintain their divergent migrations despite genetic mixing. The finding that eastern and western monarchs form one genetic population also suggests that the conservation of overwintering sites in Mexico is crucial for the protection of monarchs in both eastern and western North America.     

Phylogenetic relationships of North American nymphophiline gastropods based on mitochondrial DNA sequences

Phylogenetic relationships of 36 nymphophiline species representing 10 genera were inferred from mtCOI sequence data and compared to recent morphology-based classifications of this group. Parsimony and maximum likelihood analyses of the molecular data set suggested monophyly of the North American nymphophilines and a sister or otherwise close relationship between this fauna and a European species assigned to the subfamily. Results also supported a previously hypothesized close relationship between the predominantly freshwater nymphophilines and the brackish-water genus Hydrobia . Our analyses resolved a North American nymphophiline subclade composed of Floridobia , Nymphophilus , and Pyrgulopsis , and depicted the remaining North American genera ( Cincinnatia , Marstonia , Notogillia , Rhapinema , Spilochlamys , Stiobia ) as either a monophyletic or paraphyletic group. Two of the large North American genera ( Floridobia , Marstonia ) were supported as monophyletic groups while monophyly of Pyrgulopsis , a western North American group containing > 100 species, was equivocal. North American nymphophiline phylogeny implies that vicariance of eastern and western North American groups was followed by a secondary invasion of eastern coastal areas from the west. We attribute this to dispersal of salt-tolerant progenitors along the Gulf of Mexico coast     

Bioavailability of lead in North Sea sediments

As part of an interdisciplinary research programme, lead contents were measured in the polychaeteNephtys spp. and in the sea-urchinEchinocardium cordatum as well as in the respective sediment fractions <20 μm taken from the Dogger Bank proper and the eastern coastal North Sea. A lower lead content was generally observed in the organisms taken from the German Bight than in those from the Dogger Bank, especially from its northeastern part. It is possible to divide both areas according to the slope found in the linear regression of lead versus total organic carbon contents in sediments, which is twice as steep for the Dogger Bank as for the eastern North Sea. This criterium points to a difference in sediment quality with regard to toxic metal contamination. The sediment quality of the Dogger Bank seems to be twice as bad compared with that of the eastern North Sea. This is in good agreement with the differences found in lead contamination of the sediment-dwelling polychaetes from both areas. The results indicate that lead is primarily accumulated by food ingestion.     

Reticulate evolution in Thuja inferred from multiple gene sequences: implications for the study of biogeographical disjunction between eastern Asia and North America

The eastern Asia-North America disjunction is one of the most interesting biogeographical patterns, but its formation is still in much debate. Here nucleotide sequences of five cpDNA regions, nrDNA ITS and two low-copy nuclear genes (LEAFY, 4CL) were employed to reconstruct the phylogeny and to explore the historical biogeography of Thuja, a typical eastern Asia-North America disjunct genus. High topological discordance was observed between chloroplast and nuclear gene trees, even between different nuclear gene trees, suggesting that Thuja could have a reticulate evolutionary history due to multiple interspecific hybridization events. The eastern Asian species Thuja koraiensis might have obtained its chloroplast genome from the eastern North American species T. occidentalis by chloroplast capture, while the western North American species T. plicata is very likely to have inherited a recombinant cpDNA. Based on the phylogenetic analysis of multiple genes, DIVA-reconstruction of the distribution history, molecular clock estimation and fossil data, we inferred that Thuja could have originated from the high-latitude areas of North America in the Paleocene or earlier with subsequent expansion into eastern Asia through the Bering Land Bridge. The two eastern Asia species T. standishii and T. sutchuenensis have a sister relationship, and their split could have occurred in the Oligocene or early Miocene. In the present study, the selection of molecular markers in biogeographic studies was also discussed. Since most previous studies on the eastern Asia and North America disjunction are based on uniparentally inherited cpDNA and (or) directly sequenced nrDNA ITS data, the historical reticulate evolution in the studied groups might have been underestimated. Therefore, we suggest that multiple genes from different genomes, especially low-copy nuclear genes, be used in this research area in the future.     

Floristical and ecological characterization of the polar desert zone of Greenland

Abstract. Species composition and biomass of four plant communities were investigated in two coastal polar desert areas in eastern North Greenland, bordering the North East Water Polynya - an ice-free sea area kept open by upwelling - and compared with inland areas in North Greenland. Herb barren, the poorest type, has a species richness of 6 species/m², a cover of 0.7 %, and an aboveground biomass of 0.6 g/m² (vascular plants). The richest type, Saxifraga oppositifolia snowbed, has 10 species/m², 5.0 % cover, and 11.2 g/m² biomass. A floristic and vegetation boundary exists a few kilometres from the coast. The coastal areas bordering the North East Water Polynya had an impoverished flora and vegetation compared to areas near the ice-covered sea, possibly caused by very low summer temperatures and high frequency of clouds. A new delimitation of the polar deserts of Greenland is proposed on the basis of the number of vascular plant species, the occurrence of species with a specific inland distribution in North Greenland and the dominating life forms. At present the polar desert zone includes only areas within a zone up to ca. 15 km from the outer coast of high arctic Greenland - north of ca. 80° N. Large areas formerly classified as polar deserts in eastern North Greenland, as well as in Washington Land in western North Greenland, are excluded. New floristic data confirm that Greenland is correctly included in the Canadian province of the arctic polar deserts, whereas there is no reason for subdividing the polar deserts of the Canadian province.     

Phylogenetics and biogeography of eastern Asian-North American disjunct genus Pachysandra （Buxaceae） inferred from nucleotide sequences

Pachysandra is an eastern Asian-North American disjtunct genus with three species, two in eastern Asia （Pachysandra axillaris and Pachysandra terminalis） and one in eastern North America （Pachysandra procurnbens）. Although morphological and cytological studies suggest a close affinity of Pprocumbens with P axillaris, molecular data from nuclear and chloroplast DNA regions have provided conflicting signals. In this study, we tested previous phylogenetic hypotheses using sequences of nuclear ribosomal DNA internal transcribed spacers and chloroplast ndhF gene from multiple individuals of each of the three species. We also estimated the time of divergence between eastem Asia and eastern North America. Our results support the morphological and cytological conclusion that P procumbens is more closely related to P axillaris than to P terminalis. The estimated time of divergence of P axillaris and P procumbens was 14.6±5.5 mya, consistent with estimates from many other eastern Asian-North American disjunct genera. The migration of Pachysandra populations from eastern Asia to North America might have occurred by way of the North Atlantic land bridge.     

GENETIC DATA HINT AT A COMMON DONOR REGION FOR INVASIVE ATLANTIC AND PACIFIC POPULATIONS OF GRACILARIA VERMICULOPHYLLA (GRACILARIALES,RHODOPHYTA)1

Gracilaria vermiculophylla (Ohmi) Papenf., an agar‐producing red alga introduced from northeast Asia to Europe and North America, is often highly abundant in invaded areas. To assay its genetic diversity and identify the putative source of invasive populations, we analyzed the mitochondrial cytochrome c oxidase subunit I (cox1) gene from 312 individuals of G. vermiculophylla collected in 37 native and 32 introduced locations. A total of 19 haplotypes were detected: 17 in northeast Asia and three in Europe and eastern and western North America, with only one shared among all regions. The shared haplotype was present in all introduced populations and in ～99% of individuals in the introduced areas. This haplotype was also found at three native locations in east Korea, west Japan, and eastern Russia. Both haplotype and nucleotide diversities were extremely low in Europe and North America compared to northeast Asia. Our study indicates that the East Sea/Sea of Japan is a likely donor region of the invasive populations of G. vermiculophylla in the east and west Atlantic and the east Pacific.     

TAXONOMIC STATUS AND GEOGRAPHICAL CRANIAL VARIATION OF COMMON DOLPHINS (DELPHINUS) IN THE EASTERN NORTH ATLANTIC

The common dolphin has a widespread distribution and is relatively abundant in the temperate to subtropical waters of the eastern North Atlantic. However, it is not known whether different species, subspecies, or populations occur in this region. We examined 393 common dolphin skulls obtained from both stranded and bycaught individuals collected between 1901 and 2005. The series included skulls of 152 females and 199 males, from animals ranging in body length from 93 to 230 cm and 105 to 244 cm, respectively. The ranges of total body length, skull size, RL/ZGW ratio and maximum upper alveolar (tooth) count of common dolphins in the eastern North Atlantic overlapped with those of both short- ( D. delphis ) and long-beaked ( D. capensis ) species found off the Californian coast. However, in the absence of additional data, the common dolphin in the eastern North Atlantic is regarded here as a large form of Delphinus delphis . Sexual dimorphism and possible sex-linked characters were identified within the sample. Results of the current study indicate some population differentiation within the eastern North Atlantic, with common dolphins off Portugal showing segregation in morphometric characteristics from common dolphins in other areas.     

Molecular phylogeography of domesticated barley traces expansion of agriculture in the Old World

Barley (Hordeum vulgare ssp. vulgare) was first cultivated 10,500 years ago in the Fertile Crescent and is one of the founder crops of Eurasian agriculture. Phylogeographic analysis of five nuclear loci and morphological assessment of two traits in >250 domesticated barley accessions reveal that landraces found in South and East Asia are genetically distinct from those in Europe and North Africa. A Bayesian population structure assessment method indicates that barley accessions are subdivided into six clusters and that barley landraces from 10 different geographical regions of Eurasia and North Africa show distinct patterns of distribution across these clusters. Using haplotype frequency data, it appears that the Europe/North Africa landraces are most similar to the Near East population (F ST = 0.15) as well as to wild barley (F ST = 0.11) and are strongly differentiated from all other Asian populations (F ST = 0.34-0.74). A neighbor-joining analysis using these F ST estimates also supports a division between European, North African, and Near East barley types from more easterly Asian accessions. There is also differentiation in the presence of a naked caryopsis and spikelet row number between eastern and western barley accessions. The data support the differential migration of barley from two domestication events that led to the origin of barley--one in the Fertile Crescent and another farther east, possibly at the eastern edge of the Iranian Plateau--with European and North African barley largely originating from the former and much of Asian barley arising from the latter. This suggests that cultural diffusion or independent innovation is responsible for the expansion of agriculture to areas of South and East Asia during the Neolithic revolution.     

Phylogeny and biogeography of Aralia sect. Aralia (Araliaceae)

Aralia sect. Aralia (Araliaceae) consists of approximately eight species disjunctly distributed in Asia and North America. Phylogenetic and biogeographic analyses were conducted using sequences of the internal transcribed spacer regions of the nuclear ribosomal DNA. Aralia racemosa from eastern North America was sister to A. californica from western North America. Aralia cordata from eastern Asia did not form a species-pair relationship with the eastern North American A. racemosa. The two subspecies of A. racemosa formed a monophyletic group. Biogeographic analyses showed a close area relationship between eastern North America and western North America. The Himalayas were cladistically basal and eastern Asia was placed between the Himalayas and North America. The biogeographic analysis supported the origin of the eastern Asian and eastern North American disjunct pattern in Aralia sect. Aralia via the Bering land bridges. Comparisons with results of phylogenetic analyses of other genera suggested that (1) the floristic connection between eastern North America and western North America may be stronger than previously thought; and (2) the biogeographic patterns in the Northern Hemisphere are complex. Furthermore, a lack of correlation between sequence divergence values and phylogenetic positions was observed, suggesting the importance of a phylogenetic framework in biogeographic analyses.     

Variation in spring migration routes and breeding distribution of northern pintails Anas acuta that winter in Japan

In North America, spring migration routes and breeding distribution of northern pintails Anas acuta vary because some individuals opportunistically nest at mid‐latitudes in years when ephemeral prairie wetlands are available, whereas others regularly nest in arctic and sub‐arctic regions where wetland abundance is more constant. Less was known about migration routes and breeding distribution of pintails in East Asia. From 2007–2009 we marked 198 pintails on their wintering areas in Japan with satellite transmitters to: 1) document spring migration routes and summer distribution, 2) evaluate migratory connections and breeding season sympatry with North American pintails, and 3) determine if pintails used the same migration routes in fall as in spring. Most pintails (67%) migrated to the Kamchatka or Chukotka peninsulas in eastern Russia either directly from Japan or via Sakhalin Island, Russia. Remaining pintails primarily migrated to the Magadan region or Kolyma River Basin in eastern Russia via Sakhalin Island. The Chukotka Peninsula was the most common summer destination, with highest densities in the Anadyr Lowlands; a region also used by pintails that migrate from North America. One pintail migrated to St. Lawrence Island, Alaska, in spring and another briefly migrated to the western coast of Alaska in fall. Autumn migration routes generally mirrored spring migration although most pintails bypassed Sakhalin Island in fall. Compared to North American pintails, pintails that winter in Japan exhibited less variation in migration routes and breeding distribution, and nested at higher latitudes. In the Russian Far East there is no region with habitats comparable in extent to the ephemeral mid‐latitude wetlands of North America. Consequently, East Asian pintails mainly nest in arctic and sub‐arctic regions where annual consistency in wetlands promotes constancy in migration routes and breeding distribution. Breeding season sympatry between pintails from different continents results more from North American pintails migrating to eastern Russia than from Japanese pintails migrating to North America.     

A New Fishfly Species (Megaloptera: Corydalidae: Neohermes Banks) Discovered from North America by a Systematic Revision,with Phylogenetic and Biogeographic Implications

The taxonomy of Megaloptera from the Nearctic region is fairly well known and their faunal diversity has been largely surveyed, even in relatively remote regions. However, the evolutionary history of Nearctic Megaloptera is still poorly known with phylogenetic and biogeographic studies lacking. In this paper, we report a new fishfly species of the endemic North American genus Neohermes Banks, 1908, increasing the total number known of species to six. This new species (Neohermes inexpectatus sp. nov.) is currently known to occur only in California (USA) and is apparently confined to the Northern Coastal Range. The new species resembles the three Neohermes species from eastern North America based on the relatively small body size and the presence of female gonostyli 9. However, our phylogenetic analysis using adult morphological data recovered the new species as the sister species to the remaining Neohermes, which includes two species from western North America and three from eastern North America. According to the present interspecific phylogeny of Neohermes, with reconstructed ancestral areas, the initial divergence within the genus was found to take place in western North America, with a subsequent eastward dispersal. This likely lead to the modern distribution of Neohermes in eastern North America with the closure of the Mid-Continental Seaway, which separated western and eastern North America in the Mid-Late Cretaceous (100–80 MYA) and finally disappeared at the end of the Cretaceous (70 MYA). The uplift of the Cordilleran System probably accounted for the divergence between the eastern and two western Neohermes species.     

Evolution of the Eastern Asian and Eastern North American disjunct genus Symplocarpus (Araceae): insights from chloroplast DNA restriction site data

Chloroplast DNA (cpDNA) variation was surveyed with 20 restriction endonucleases for the eastern Asian and eastern North American disjunct genus Symplocarpus (Araceae). The cpDNA phylogeny reveals a sister group relationship between S. foetidus from eastern North America and S. renifolius from eastern Asia. The cpDNA divergence between the two intercontinental sister species is 0.61%, which suggests an estimated divergence time of 6.1 million years ago during the late Miocene. The Bering land bridge hypothesis is compatible with the estimated time of divergence for the migration of Symplocarpus between eastern Asia and North America. Furthermore, a single origin of the exothermic spadices in Symplocarpus is suggested by the phylogeny. The cpDNA data also provide independent support for the recognition of three species within the genus.     

Phytogeographical and community similarities of alpine tundras of Changbaishan Summit,China, and Indian Peaks,USA

Abstract. We compared the diversity, phytogeography, and plant communities in two mid-latitude alpine tundras with comparable aerial and elevational extents: Changbaishan Summit in eastern Asia and Indian Peaks in western North America. Despite wide separation, the two areas shared 72 species. In all, 43% of the species on Changbaishan Summit are also distributed in the alpine zones of western North America, while 22% of the species on Indian Peaks are also distributed in the alpine zones of eastern Asia. Almost all the shared species also occur in the Beringian region. Phytogeographical profiles of species and genera showed that 69% of species and over 90% of genera in both alpine tundras belong to the three phytogeographical categories: cosmopolitan, circumpolar, and Asian-North American. We attributed the current floristic relationship between these widely separated areas to the periodic past land connection between the two continents during the Tertiary and Pleistocene. Indian Peaks has a closer floristic relationship with the Arctic tundra than does Changbaishan Summit. Indian Peaks also has 45% higher species richness and lower vegetation cover than Changbaishan Summit. Plant communities from the two areas were completely separated in the two-way indicator species analysis and non-metric multidimensional scaling on floristic data at both species and generic levels, whereas ordination of communities by soil data produced a greater overlap. The plant communities on Changbaishan Summit in general have lower alpha diversity, higher beta diversity (lower between-community floristic similarity), and more rare species than does Indian Peaks. Mosaic diversity does not differ in the two alpine tundras, although the analysis suggests that Changbaishan Summit communities are more widely spaced on gradients than the Indian Peaks communities.     

Phylogenetics and biogeography of eastern Asian–North American disjunct genus Pachysandra (Buxaceae) inferred from nucleotide sequences

Abstract Pachysandra is an eastern Asian–North American disjunct genus with three species, two in eastern Asia (Pachysandra axillaris and Pachysandra terminalis) and one in eastern North America (Pachysandra procumbens). Although morphological and cytological studies suggest a close affinity of P. procumbens with P. axillaris, molecular data from nuclear and chloroplast DNA regions have provided conflicting signals. In this study, we tested previous phylogenetic hypotheses using sequences of nuclear ribosomal DNA internal transcribed spacers and chloroplast ndhF gene from multiple individuals of each of the three species. We also estimated the time of divergence between eastern Asia and eastern North America. Our results support the morphological and cytological conclusion that P. procumbens is more closely related to P. axillaris than to P. terminalis. The estimated time of divergence of P. axillaris and P. procumbens was 14.6±5.5 mya, consistent with estimates from many other eastern Asian–North American disjunct genera. The migration of Pachysandra populations from eastern Asia to North America might have occurred by way of the North Atlantic land bridge.     

A review of vital rates and cause‐specific mortality of elk Cervus elaphus populations in eastern North America

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Comparison of the post-Chernobyl 137Cs contamination of mushrooms from eastern Europe, Sweden, and North America.

A comparison was made of 134Cs and 137Cs contamination in fungi from eastern Europe and eastern North America. Mean activities of 25 Ukrainian, 6 Swedish, and 10 North American collections were 4,660, 9,750, and 205 Bq/kg (dry weight), respectively. Additional measurements were made on samples from the Moscow, southern Belarus, and Yugoslavia/Bulgaria regions. Activity values were found to vary by several orders of magnitude within all geographic areas, even for the same mushroom species. Significantly higher specific activities were observed in mycorrhizal species than in saprophytic and parasitic fungi. Unfortunately, many of the European mycorrhizal species considered as prized edibles contained unacceptably high levels of 137Cs (> 1,000 Bq/kg [dry weight]) and should be used sparingly as food. By contrast, no mushrooms collected in Ontario or northern Michigan exceeded 1,000 Bq of 137Cs per kg (dry weight). The excessive 137Cs contamination was evident in mushrooms from areas that had substantial fallout from the 1986 accident in reactor 4 at the Chernobyl nuclear power station. However, observations suggest that about 20% of the 137Cs in eastern Europe (Moscow area, Belarus, and Ukraine) is of non-Chernobyl origin.